Testosterone Administration Decreases Generosity in the Ultimatum Game

Paul J. Zak1*, Robert Kurzban2, Sheila Ahmadi3, Ronald S. Swerdloff4, Jang Park1, Levan Efremidze1, Karen Redwine1, Karla Morgan5, William Matzner1
1 Center for Neuroeconomics Studies, Claremont Graduate University, Claremont, California, United States of America, 2 University of Pennsylvania, Philadelphia, Pennsylvania, United States of America, 3 Department of Endocrinology, University of California Los Angeles, Los Angeles, California, United States of America, 4 Division of Endocrinology, Harbor University of California Los Angeles Medical Center, Torrance, California, United States of America, 5 Department of Economics, Whitworth University, Spokane, Washington, United States of America

How do human beings decide when to be selfish or selfless? In this study, we gave testosterone to 25 men to establish its impact on prosocial behaviors in a double-blind within-subjects design. We also confirmed participants’ testosterone levels before and after treatment through blood draws. Using the Ultimatum Game from behavioral economics, we find that men with artificially raised T, compared to themselves on placebo, were 27% less generous towards strangers with money they controlled (95% CI placebo: (1.70, 2.72); 95% CI T: (.98, 2.30)). This effect scales with a man’s level of total-, free-, and dihydrotestosterone (DHT). Men in the lowest decile of DHT were 560% more generous than men in the highest decile of DHT. We also found that men with elevated testosterone were more likely to use their own money punish those who were ungenerous toward them. Our results continue to hold after controlling for altruism. We conclude that elevated testosterone causes men to behave antisocially.
Citation: Zak PJ, Kurzban R, Ahmadi S, Swerdloff RS, Park J, et al. (2009) Testosterone Administration Decreases Generosity in the Ultimatum Game. PLoS ONE 4(12): e8330. doi:10.1371/journal.pone.0008330 ´ Aleman, University of Groningen, The Netherlands Editor: Andre Received June 9, 2009; Accepted November 26, 2009; Published December 16, 2009 Copyright: ß 2009 Zak et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: The authors thank the MacArthur Foundation Research Network on The Nature and Origin of Preferences and the John Templeton Foundation for support. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: paul@pauljzak.com

Human beings are both prosocial and self-serving, often exhibiting both behaviors in a short period of time. The neurologic foundations for prosociality are just beginning to be examined [1–4], but the mechanisms that cause a shift from selfless to selfish have not been characterized. There is an extensive literature associating male aggressive and antisocial behaviors with elevated testosterone (T) [5,6]. Yet, T is not the most obvious candidate promoting selfishness; the recent multi-billion dollar donations to charity by Bill Gates and Warren Buffett reveal that males with significant resources may be generous. Studies in monkeys show that when beta males become alphas, both T and serotonin rise while cortisol falls [7,8]. Alpha males have been observed sharing resources, but this is typically strategic, for example, to sustain a supporting coalition [9,10]. Yet, alpha males, unlike lower ranking members of a social group, may have less need to be generous towards others. Correlational studies of salivary T in humans have found that high T males are more likely to have physical altercations, divorce more often, spend less time with their children, engage in competitions of all types, have more sexual partners, face learning disabilities, and lose their jobs more often [11,12] suggesting that high T men may behave differently than other men. A recent study found that high T males are more likely to reject stingy offers in the Ultimatum Game [13], but whether high T is the cause or the effect of a low offer is unclear; low offers in a related task called the trust game have been
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associated with a rise in serum dihydrotestosterone [14]. More generally, high T males appear to be more aggressive and less prosocial [15]. These correlations should be viewed with caution as T is highly dependent on a variety of environmental conditions [11]. For example, winning a chess match will raise T, and watching one’s team lose a soccer game on TV will cause T to fall [16,17]. The inability to control experimental subjects’ behaviors before they enter the lab, and the high degree of variability in basal T indicate that correlational studies can only be considered provisional findings [18]. In addition, salivary testosterone assays, while convenient, have measurement problems, including the effect of foreign substances such as gum to facilitate salivation and contamination with blood due to microtrauma. Further, there is only a moderate correlation between T measured in saliva and blood serum [19]. Critiquing correlational studies of T and behavior, O’Carroll wrote that ‘‘Definitive evidence is likely to come from placebocontrolled, double-blind experiments in which circulating T levels are manipulated and appropriately reliable and sensitive assays of behaviour are taken.’’ [18]. Manipulating T produces direct causal evidence directly relating to T to behavior, and this is precisely the approach we take here. Yet, hormone manipulation is rare in the nonclinical literature. Studies that infused moderate supraphysiologic doses of T into eugonadal males have found little effect on anger or mood [20–22]; mood effects occur only for very high doses of T. A very small study (N = 6) showed that men who had
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participants arrived at 4 pm and were interviewed by a licensed medical doctor (S. medications that interact with T. participants were led to a semi-private booth. Within a dyad.0008330. Only male participants were recruited because the US Food and Drug Administration has only approved testosterone treatment for men. both DMs received extensive and identical instructions regarding how their decisions and those of the other DM in the dyad would affect how much money each could make. All participants made proposals as DM1s and as DM2s were asked to state their minimum acceptable proposal to elicit their punishment thresholds. the Ultimatum game (UG) and the Dictator Game (DG). Participants appeared to understand this because there was variation in UG choices across rounds (average within-subjects SD of: proposals $0. In all tasks. and the sample was ethnically diverse (Asian 44%.) for possible contraindications for T administration.Testosterone and Selfishness their T raised for six weeks. The subgame perfect equilibrium is for DM1 to offer $1 and for DM2 to accept this. Other/no data 12%). All decisions were made by computer in partitioned stations and without communicating to others in the experiment. There is agreement in experimental economics that using the strategy method with payments produces very similar data to real-time dyadic matching. Participants were fully instructed that all decisions were ‘‘live’’ in that after making decisions. and men were likely to be more reactive behaviorally to its effects [25]. Both DMs were fully and identically instructed in this task.pone. the roles were randomized to determine earnings.org 2 other people. 29).A. The experiment was approved by the Institutional Review Boards of UCLA and Claremont Graduate University. In every session. punishment threshold $1. an Anger Inventory [4].8 years old (SD = 2. and drug or alcohol abuse. and were given a colorless hydroalcoholic gel containing either 10 g of AndrogelH (1% testosterone gel) or an inert substance. and generosity $1. The questionnaires measured demographic. After the second blood draw. payment was determined by randomly assigning each person to the role of DM1 or DM2 for each decision. asked to remove their shirts. participants were randomly assigned by computer to dyads. that involve money and PLoS ONE | www. These tasks also allowed us to measure the incidence of punishment of those who violate an implicit social norm of generosity.2). Following published pharmacokinetics [26] on peak levels of T. Session sizes varied from four to eight participants. All participants were asked to make proposals as DM1s and to identify their minimum their acceptable offers as DM2s. which measures dispositional empathy. In the UG. On debriefing. cross-over study to examine the effects of T on social behaviors. participants completed questionnaires by computer using a random alphanumeric code as their only identifier.68. Exclusion criteria included significant medical or psychiatric illness. 1). Hispanic 8%. All participants gave written informed consent for the study. Decision-Maker 1 (DM1) is endowed with $10 and DM2 has no endowment. In the Ultimatum Game.19. there was a decision-maker 1 (DM1) and decision-maker 2 (DM2). After making decisions as DM1 and DM2. The UG and DG are standard tasks in experimental economics and neutral language in the instructions was used throughout. doi:10. Twenty-five participants completed the entire experiment and are included in our analyses. At the end of the experiment. After medical screening and consent. we manipulated T in healthy eugonadal men in a double-blind. DM2 could either accept the proposal and then the money would be paid. Accepted proposals are paid to both DMs while rejected proposals cause both DMs to receive nothing. For every session. Participants were instructed and observed spreading the gel on their shoulders and upper back following the AndrogelH instructions. or he could reject the proposal and both DM2s would get nothing (Fig. Using a neuroeconomics paradigm [24]. The blood draw for the 8am session established how much higher participants’ T levels were after AndrogelH administration. No adverse events were reported. We hypothesized that participants given T would be more likely to punish those making ungenerous monetary offers to them. participants had 28 ml of blood draw from an antecubital vein.g001 December 2009 | Volume 4 | Issue 12 | e8330 . we are able to perform within-subjects comparisons of behavior. social. and the Personal Reaction Inventory (PRI)[31] that measures social behaviors. Next. Participants next made decisions in two tasks. The mean age of participants was 20. In the present study. to answer survey questions. DM1 was prompted by computer to propose a split of this money to DM2. By using the ‘‘strategy method’’ in which participants make both proposals at DM1s and state their punishment threshold as DM2s [32]. DM1 was endowed with $10 while DM2 had nothing. 28. Affective Intensity Measure that addresses emotional responses (AIM) [30]. the Interpersonal Reactivity Index (IRI. participants returned to the lab 16 hours after administration for a second blood draw. DM1 proposes a split of his endowment to DM2 that DM2 can either accept or reject. participants made a set of decisions involving money. and psychological traits. Figure 1. approximately one-half of the participants received testosterone and the other half were given the placebo. compared to themselves on placebo. participants reported that they did not know which substance they had been given. After instruction. Materials and Methods Forty-eight male students were recruited for this double-blind cross-over experiment. In both the UG and DG. These included Experiences in Close Relationships-Revised (ECR-R) [27] that measures attachment styles. and make a series of decisions involving money. Caucasian 36%. with study phases (testosterone or placebo) separated by six to 12 weeks depending on which sessions participants were in. DM1 to DM2 pairings would be made that determined their earnings. were more likely to respond in kind to a perceived provocation (though actually fictitious) in which they were made to lose a small amount of money by another person [23].plosone.1371/journal. We hypothesized that T would cause men to behave less generously towards strangers.55).

Free T (Inter-assay: 5.79. ng/ml (SD 3. Biomarkers Core using kits from Diagnostic Systems Laboratories (Webster. Values of total T.60% at 23. T infusion did affect the amount of negative generosity (proposals. DHT (Inter-assay: 7. In Western countries. 809.30 pg/ml. All of these changes were greater than zero for p. In addition. two-tailed paired t-test.68 ng/ml. free T.32 pg/ml or a 170% change from baseline.1E-6 (two-tailed paired t-tests. n = 2.84. free T: 15. Placebo: $2.63.). 4. total T was 60% higher. free T: 14.2 pg/ml (SD = 228. We next assessed whether T was higher after 10 g of AndrogelH treatment.93). To confirm our results. we established that for those receiving the placebo that T levels did not rise overnight. and generosity. Intra-assay: 7. the large sample size and paired data with a central moment and kurtosis indicate that t-tests are appropriate for the analysis [36–38]. All assays were performed by Yerkes Figure 2. Placebo: $2. Proposals of exactly the minimum acceptable amount are not generous because they do not demonstrate ‘‘liberality in giving’’ or offering more than another person expects or needs. Total T (Inter-assay: 1.48). Participants made decisions in the UG and DG four times with random rematching to other DMs each round.63) paired t-test p = . one-tailed paired t-test. 10. At the end of the experiment.9% rejections for participants on placebo (p = . Sixteen hours after AndrogelH treatment.9). Average T levels prior to AndrogelH treatment were total T: 4. Consistent with our primary hypothesis. the smallest change in total T was 0.85.03 pg/ml. Participants were instructed that they would make four one-shot decisions.05. There was no deception of any kind.2 pg/ml (SD . and DHT: 753.86 ng/ml. n = 6.Testosterone and Selfishness Following a related study.046.99). 1.1 ng/ml (SD 3. In this task. Behavior Average DM1 proposals in the UG were 9% lower for men on T compared to themselves on placebo (T: $4. The UG can also be used to measure the willingness by individuals to engage in costly punishment of stingy offers or for violations of implicit sharing norms.rejection threshold) by men on T compared to themselves on placebo was 27% lower (T: $1.31 pg/ml or a 7% change. two-tailed t test).6% of participants on AndrogelH rejecting their own proposals compared to 2. p = .08. DHT: 704.4 ng/ml (SD 3. 4. N = 200. 736.34). we ran a random-effects GLM of generosity and a T indicator variable for DM1 offers.39). average T values before infusion and 16 hours later were unchanged (total T: 4. free. because T was elevated in every participant given AndrogelH compared to himself. Fig. participants were paid their earnings privately by a lab administrator.1371/journal. Intra-assay: 1. the UG was used to measure generosity [33]. After instruction. The blue bar is basal T and the blue plus the red bar is the post-treatment T value (and SE bars).001). n = 2). all differences p. with 9. N = 200. T in every treated subject was higher than baseline. paired t-test p = .15. A high minimum acceptable offer therefore punishes DM1 for stingy offer but at a cost to DM2. TX). We assayed total. offers less than 30% of DM1’s endowment are nearly always rejected [32].92.57. p = . the DM2 rejection threshold was 5% higher on T versus placebo though the difference was not significant (T: $3. participants were also randomly put into dyads in which DM1 had $10 and DM2 had zero.1E-6.87 pg/ml. The highest level of total T after treatment was 10. For men receiving placebo. doi:10. DM2 rejection threshold. Although the data are not normally distributed (Shapiro-Wilk test p = . n = 4. n = 4. Stingy offers to DM2s in the UG have been shown to provoke anterior insula activity [34] suggesting that low offers are rejected due to a sense of disgust.92).04 pg/ml. Placebo: $5.32% at 118. DM2 had no choice to make.08). and dihydrotestosterone (DHT) to fully characterize the androgenic state of participants. Fig. one-tailed paired t-test.6).0 pg/ml (SD .3 pg/ml (SD . Money transferred in the DG is thought to measure altruism [32]. 3). at 0.org 3 December 2009 | Volume 4 | Issue 12 | e8330 . Our basic findings showing Results Testosterone First.g002 PLoS ONE | www.67 ng/ml. 2.14% at 5. free T was 97% higher.035. p = .3 pg/ml (SD 413.1 pg/ml (SD = 267.55% at 3. This approach was used to expose fewer participants to the effects of drugs following a protocol we have previously used with oxytocin infusion [35]. N = 200. The DG was included as a control.17% at 624.rejection threshold).pone. Intraassay: 6. and DHT was 128% higher (SDs. N = 50).3%. paired t-test p = .plosone. and DHT before and after AndrogelH treatment. DM1 was asked to make a unilateral offer of some of his endowment to the DM2 in the dyad. generosity (proposals . no one was excluded from the analyses.64. At the same time.001).61). CVs for assays where within acceptable ranges. A generous offer is defined as the difference between the DM1 proposal and the participant’s own minimum acceptable offer as DM2.95% at 0.52).0008330.

9%). or DHT separately) and a binary order indicator again the negative relationship between T and generosity continues to be significant (free T. generosity.0. the threshold to initiate costly punishment for those who are less generous towards them increases with T levels. Anger (p = . Nearly identical results obtain when those receiving placebo in the first stage are analyzed.1371/journal.pone.2284. and DHT (r = 0.009. we found that men whose T was elevated due to AndrogelH continued to be less generous (total T: b = 2. p = 0. generosity: coeff. p = 0.27. IRI (p = .57.0004). rejection was 30% ($0.017.17).001. see Fig. DHT: p = . we analyzed the survey responses of participants to examine their affective states on and off AndrogelH. we controlled for altruism and again examined the effect of testosterone on generosity in a leastsquares regression. p = 0. twotailed paired t-test. p = 0. This indicates that participants became more generous during the course of the experiment. p = 0. free T (r = 0. PRI (p = . avoidant attachment p = .076. = . We also had participants make decisions in the DG in order to dissociate generosity and altruism [33]. total. while generosity is giving more than the other needs. as measured by offers in the DG. p = .0011). only for those receiving AndrogelH in the first phase.001.007. DHT: b = 2. 4B. This indicates that temperament and mood were stable throughout the experiment.UG punishment threshold) by participants on placebo was $2.plosone. Testing all three measures of T (free.0. generosity and punishment maintained significance controlling for the order of AndrogelH administration. No parametric relationship between DG offers and any measure of T was found.52.Testosterone and Selfishness Figure 3. Interestingly.0068). we tested if behavior differed when a participant received AndrogelH or placebo on the first phase of the experiment as compared to the second phase.84). More participants on AndrogelH relative to placebo showed negative generosity by setting a punishment threshold above than their own offer to DM2 (9.057.25. rejection threshold: coeff.3063. and DHT (r = 20. the latter being a subset of the former. and DHT). We found that those who received AndrogelH in phase one were 78% ($1.1937. all two-tailed t-tests). free T: b = 20.0.05. Running a least squares regression on generosity. Significant correlations were also found for the rejection threshold and total T (r = 0.81).44. p = . The parametric relationship between T levels and reduced generosity continued to maintain significance when DG offers were included (total T: b = 20. p = 0. total T: p = . p values for possible differences are: ECR-R (overall.0001.0008330.88) higher (two-tailed t-test.003. As a result. Because of these effects. p = 0. N = 88). December 2009 | Volume 4 | Issue 12 | e8330 . 4A. p = 0.11.6% vs. A similar effect was found on the rejection threshold.001). was not different for those on AndrogelH compared to placebo (T: $3.013. and punishment. DHT: b = 20.g003 that T makes men less generous continue to hold (DM1 offers: coeff. we examined if there was a parametric relationship between T. Behavioral studies of strategic economic games have found learning effects from repeat play [32].038.01) for participants getting T in the first phase compared to those given placebo. p = 0. Participants on AndrogelH were marginally more emotionally labile (AIM.64). the change in T (total. Discussion Our primary finding is that manipulating T in men causes them to be 27% less generous in the UG then themselves at baseline.009.048. anxious attachment p = .400. p = . p = 0. p = 0. p = 0.0060). p = 0. p = 0.46. we found that greater T was associated with less generosity and an increased desire to punish those making stingy offers. p = .001. DHT: b = 0.55. Placebo: $3.15 compared to $1. 2.69). We ran the same analysis for the rejection threshold and PLoS ONE | www. free. p = 0.68) less generous than participants on placebo (two-tailed t-test.035. change in T values.0306). Across the two sessions.1908.57 when the same individuals were given AndrogelH.001. Generosity (UG offer . bars in graph are SEs). a 27% reduction (p = . free T: b = .001.07). see Fig. Lastly. p = . p = .03.56. free T: b = 2.001). p = .34. = 2. = 2.86). two-tailed t test.org 4 again found that punishment of those who were not generous increased with change in T levels (total T: b = . Including the entire N = 200 data set in a least squares regression for generosity. free T (r = 20. Next.0001). For generosity we found highly significant correlations with total T (r = 20. p = . we tested whether the parametric relationship between T. N = 200 for this and subsequent tests. N = 88). Altruism is defined as giving to help another. Altruism. DM2s showed a lower likelihood of punishing stingy offers. p = 0. doi:10.001. Because differences in altruism might impact generosity. Using paired two-tailed t-tests.1529.

(A) The reduction of generosity for those on AndrogelH positively scales with levels of total T.3063. Our results are not due to T making men December 2009 | Volume 4 | Issue 12 | e8330 . p = 0. doi:10.1371/journal. This increase in negative generosity between conditions suggests that T infusion interfered with participants’ ability to understand others’ behaviors since rejections of DM1 proposals do not earn participants any money.55 for men in the highest decile of DHT (85% lower). Men in the lowest decile of DHT had average generosity of $3.15 compared to a punishment threshold of $4. participants on AndrogelH were more than twice as likely to have exhibited negative generosity (rejection threshold exceeding proposed split) compared to themselves on placebo.org 5 through blood draws. free T and DHT. These results are credible because T was directly manipulated. generosity for those on AndrogelH are shown in red. the relationship for DHT is shown (r = 0.2284.00 for men in the top decile of DHT (86% higher). free T and DHT.pone.65 compared to generosity of $0.g004 Indeed.plosone. Our findings suggest that men with naturally high T levels would be expected to be more selfish and also more likely to punish others for violations of social norms. the relationship for DHT is shown (r = 20. Scale effect of T on generosity. the effects of T on generosity and punishment scale with a man’s T levels. Men in the lowest decile of DHT had average punishment threshold of $2. consistent with many correlational studies using retrospective reporting of behaviors and salivary T measures (11).0001). p = 0. (B) The punishment threshold also scales with a man’s level of total T.Testosterone and Selfishness Figure 4. Further.0008330. and the comparisons are within-subjects. and the change in T was documented PLoS ONE | www.0011). T levels and generosity for those on placebo are shown in blue.

O’Carroll RE (1998) Placebo-controlled manipulations of testosterone levels and dominance. Am J Primat 42(1): 25– 39. Harris JH. Starzyk KB. Our previous research showed that blood draws do not affect adrenocorticotropic hormone (ACTH) levels [46]. Performed the experiments: PZ RK SA JP KM. 5. Brennan KA (2000) An item response theory analysis of self-report measures of adult attachment. Primates 24(3): 318–336. 11. T administration may have influenced the functioning of the hypothalamic-pituitary-adrenal (HPA) axis as there is significant cross-talk between these systems [45]. speculative. Raleigh MJ. Shirtcliff EA. Acknowledgments We acknowledge the encouragement and advice from Herb Gintis. Agg Beh 22: 321–331. This may explain a variety of gender differences in seeking to enforce rules of conduct. 18. Booth A (1998) Testosterone and dominance in men. at this point. Singer T (2007) The neuronal basis of empathy and fairness. et al. By administering T. Book AS. Bernhardt PC. Physiol Behav 65: 59–62. and their generosity in the UG scaled positively with their subjective empathy ratings. 7. 25. our findings can provide insights about the origins of selfish and violent behaviors ranging from reckless driving. Those with reduced serotonin rejected approximately 85% of highly unfair offers (20% of DM1 endowment) compared to an approximately 70% rejection rate for these offers for placebo participants. Pope HG. Nishida T (1983) Alpha status and agonistic alliance in wild chimpanzees (Pan troglodytes schweinfurthii). Nature 415: 125–128. Psychoneuroendocrin 29: 1229–1240. The impact of T on OT in the UG is. Neuron 60: 409–411. Zak PJ (2008) The neurobiology of trust. 31. Yet seven patients with ventromedial prefrontal lesions who played the UG as DM2s rejected 20% of endowment offers 74% of the time. 17. Diener E. In a related study. Am Ec Rev: Papers Proc 95(2): 360–363. Watts DP (2001) Why do chimpanzees hunt and share meat? Animal Beh 61: 915–924. Beh Brain Sci 21(3): 382–383. Storer TW. 3. In that study. Holman T. This suggests that the T administration increasing the punishment threshold is not due to HPA axis effects. 16. What we have found is that T appears to play a role inducing men to change from being selfless to being selfish. Burnham TC (2007) High-testosterone men reject low ultimatum game offers. & lovers: testosterone and behavior. References 1. 27. Zak PJ. Author Contributions Conceived and designed the experiments: PZ RK RSS WM. Snyder M (1987) Public appearances. then a high rejection threshold can be considered a prosocial behavior at odds with the stinginess high T males exhibited in proposing splits in the UG. et al. J Clinical Endocrin Metab 85(12): 4500–4510. J Personality Soc Psych 78: 350–365. Mitani JC.Testosterone and Selfishness more impulsive. Wang C. 4. 13. to watching or engaging in sporting events. Ellingrod V (1999) Psychosexual effects of three doses of testosterone cycling in normal men. Estela Hopenhayn. Cunningham G. Clevenger B. Because T responds to environmental conditions. rogues. Josephs RH (2006) Testosterone change after losing predicts the decision to compete again. Sapolsky RM (1997) Styles of male social behavior and their endocrine correlates among low-ranking baboons. Dabbs JM Jr. Kurzban R (2005) The neuroeconomics of distrust: Sex differences in behavior and physiology. 12. New York: McGraw-Hill. JSAS Catalog Select Doc Psych 10: 85. Yuwiler A (1984) Social and environmental influences on blood serotonin concentrations in monkeys. A recent paper measuring salivary T finds that high T males were more patient in waiting for rewards that were promised in the future [39]. Horm Beh 50: 684–669. generosity. David Levine.com/anger. private realities. from http://www. Dabbs JM. Wang C. Beh Brain Sci 21: 353–397. McGuire MT. Lutter CD (1998) Testosterone changes during vicarious experiences of winning and losing among fans at sporting events. Philos Trans Royal Soc B 359: 1737–1748. Swerdloff RS. Arch Gen Psychiatry 41(4): 405–410. J Clinical Endocrin Metab 81(10): 3754–3758. Yates WR.42]. Biol Psychiatry 45: 254–260. while healthy controls rejected 50% of these [49]. The parametric relationship we found between T levels. Davis MH (1983) Measuring individual differences in empathy: evidence for a multidimensional approach.org 6 December 2009 | Volume 4 | Issue 12 | e8330 . 21. Berman N. Virgin CE. we may have inhibited OT binding and reduced empathy for the other person in the dyad. Emmons RA (1986) Affect intensity and reactions to daily life events. 10. 19. Gorski R (1992) Effect of androgens on the brain and other organs during development and aging. 9. Swerdloff RS. Mehta PH.plosone. MacIndoe J. Gould RW. Quinsey VL (2001) The relationship between testosterone and aggression: a meta-analysis. 15. Kouri EM. Tricker R. Sci Am 298: 88–95. Proc Royal Soc B 274: 2327–2330. Wrote the paper: PZ RK RSS WM. Iranmanesh A. 29. Davis MH (1980) A multidimensional approach to individual differences in empathy. J Personality Soc Psych 44: 113–126. Borja K. Agg Violent Beh 6: 579–599. and an Associated Editor of this journal. those given 40 IU of OT were 80% more generous than participants on placebo. These findings can be compared to a study of generosity in the UG in which the neurohormone oxytocin (OT) was manipulated in men through intranasal infusion. Fielden JA.headyinjury. Oliva PS (1995) Increased aggressive responding in male volunteers following the administration of gradually increasing doses of testosterone cypionate. Bowles S. Kivlighan KT. New York: Freeman and Company. (1996) The effects of supraphysiological doses of testosterone on angry behavior in healthy eugonadal men-A clinical research center study. Fraley RC. males and females who were primed with an empathy-inducing video had a spike in plasma OT. 14. 23. 2. Casaburi R. 8. Matzner W T. 6. Drug Alcohol Depend 40: 73–79. (2000) Long-term pharmacokinetics of transdermal testosterone gel in hypogonadal men. Retrieved August 29. Waller NG. Brammer GL. If rejections of stingy offers is an effort to punish violators of sharing norms at a cost to oneself. This suggests that generosity is driven by feelings of empathy [40]. and no effect was found on the punishment threshold [33]. Hines M. Related research used tryptophan depletion to reduce serotonin levels and then had participants make decisions as DM2s in the UG. 30. Schwartz EB (2004) The ‘‘trouble’’ with salivary testosterone. Churchland PS (2008) The impact of neuroscience on philosophy. Brain Injury Resource Center 1998 Anger Inventory. Jackson DN (1996) Salivary testosterone and self-report aggressive and pro-social personal characteristics in men and women. to soldiers fighting in war. Dabbs MG (2000) Heroes. Hudson JI (2000) Effects of supraphysiologic doses of testosterone on mood and aggression in normal men: A randomized controlled trial. 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